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J OURNAL OF 

The Lepidopterists’ Society 



Volume 35 



1981 



Number 1 



Journal of the Lepidopterists’ Society 
35(1), 1981, 1-21 

HYBRIDIZATION OF SATURNIA MENDOCINO AND S. 
WALTERORUM, AND PHYLOGENETIC NOTES ON 
SATURNIA AND AGAPEMA (SATURNHDAE) 

Paul M. Tuskes 

1444 Henry St, Berkeley, California 94709 
AND 

Michael M. Collins 

Department of Zoology, University of California, Davis, California 95616 

ABSTRACT. The taxonomic relationship of Saturnia walterorum and S. mendo- 
cino is discussed in terms of laboratory hybrids, larval morphology, comparative life 
history, and phonetics of intermediate populations. Results indicate that these moths 
represent different taxa, but are best described as semispecies. The two taxa freely 
interbreed in the laboratory. Hybrid Fj females with walterorum as the female parent 
had normal fertility and fecundity; females from the reciprocal cross were viable but 
sterile. The two taxa are very similar morphologically and differ mainly in the dimor- 
phic female of walterorum. Populations in the southern California Coast Ranges may 
represent intergrades. A discussion of the phylogeny of the endemic California Satur- 
nia and the closely related Agapema stresses the coevolution of these moths and their 
sclerophyllous host plants in response to historic climatic changes. 



In the New World, the genus Saturnia consists of three species 
whose distribution is centered in California. The three species, Sa- 
turnia walterorum Hogue & Johnson, S. mendocino Behrens, and S. 
alhofasciata (Johnson), exhibit life history characteristics which are 
adaptations to utilizing sclerophyllous food plants in a Mediterranean 
climate of winter rains and summer drought. The localized popula- 
tions of these moths, their rapid and erratic flight, and the rugged 
terrain they inhabit, make them difficult to collect. As a result, com- 
plete and accurate life history descriptions are lacking for all three 
species. In this paper we examine the taxonomic relationship of S. 
walterorum and S. mendocino in terms of laboratory hybrids, com- 
parative life histories, larval and adult phenotypes, and phenetics of 



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Journal of the Lepidoptewsts’ Society 



geographically intermediate populations. We extend our discussion 
to include a proposed phylogenetic scheme which includes the 
closely related genus Agapema, based on biogeographic and geofloral 
data. 



Methods 

In an effort to understand the genetic relationship between Satur- 
nia walterorum and S. mendocino, we began hybridization studies in 
1974. Our walterorum stock was from Dictionary Hill, Spring Valley, 
San Diego Co., Calif., while the mendocino used in the study came 
from Thompson Canyon, near Lake Berryessa, Yolo Co., California. 
A large series of wild and reared adults from each location was ex- 
amined, and characters which represent diagnostic differences be- 
tween the two species were sought. We selected six characters which 
could be either measured or scored as to presence or absence: fore- 
wing length, dorsal forewing and hindwing discal eyespot length, the 
ratio in length between the forewing and hindwing discal eyespot, 
the presence or absence in males of a white apical patch of scales on 
the ventral surface of the forewing, and the presence or absence of a 
bold submarginal black band on the dorsal forewing surface of fe- 
males. 

Larvae secured during the study were reared outdoors in screen 
cages after the first instar on fresh branches of Arctostaphylos spp. 
maintained in water. Most larvae pupated by June, and began to 
emerge the following year during February or March. Cage matings 
were easily obtained, usually within minutes after the female began 
''calling."" Mated females oviposited readily in paper bags or other 
containers in the absence of food plant. 

The fecundity of each female was determined by measuring the 
number, size, and batch weight of the eggs. The degree of fertility 
was based on the number of eggs which hatched within each batch. 
In the field, males were obtained by means of funnel traps or wire 
cages each containing a virgin female. In this way we were able to 
sample populations more efficiently than searching for larvae or 
adults. 



Results 

In order to interpret the phenotypes of hybrid specimens we esti- 
mated the range of variation in the parental populations. The right 
fore wing length of male walterorum averaged just over 11 percent 
greater than that of male mendocino, while that of female walterorum 
was 18 percent greater than female mendocino (Table 1). A t-test 
indicated that the difference in wing length between the two species 



Table 1. Phenotypic data for S. walterorum, S. mendocino and their hybrids. 



Volume 35, Number 1 



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Measurements in mm with S.D. given below. 



4 



Journal of the Lepidopterists’ Society 



Table 2. Fecundity and fertility of S. walterorum, S. mendocino and their hybrids. 



Ova length 

No. ova Avg. ova % 

Cross laid mm S.D. wt. g hatch 



a. 9 walterorum 

b. 9 mendocino 

c. 6 walterorum x 9 mendocino = Fia 

d. 6 Fja hybrid x 9 mendocino 

e. d Fla hybrid x 9 walterorum 

f. d Fla hybrid x 9 Fia hybrid 

g. d walterorum x 9 Fia hybrid 

h. d mendocino x 9 Fia hybrid 

i. d mendocino X 9 walterorum = Fn, 

j. d Fib hybrid x 9 Fib hybrid 

k. (d Fla hybrid x 9 walterorum)2 
(d Fla hybrid x 9 mendocino)2 



114 


2.44 


0.06 


0.0035 


97.0 


83 


2.47 


0.07 


0.0037 


96.4 


77 


2.47 


0.07 


0.0037 


97.5 


73 


2.47 


0.07 


0.0037 


87.8 


137 


2.24 


0.07 


0.0034 


91.5 


33 


1.79 


0.69 


0.0025 


3.3 


52 


1.75 


0.68 


0.0017 


0.0 


44 


1.92 


0.18 


0.0022 


0.0 


130 


2.47 


0.05 


0.0036 


96.0 


96 


2.14 


0.01 


0.0036 


98.0 


73 


2.10 


0.09 


0.0037 


96.7 


75 


2.21 


0.16 


0.0032 


0.0 



and both sexes is significant (P < .05). The eyespots on the dorsal fore- 
wing and hindwing are about twice as large in walterorum as in 
mendocino. In walterorum, the forewing eyespot is larger than the 
hindwing eyespot, while in mendocino the opposite is true. Thus the 
ratio of the forewing to hindwing eyespot is greater than 1 in walter- 
orum, and less than I in mendocino. This character was scored qual- 
itatively and not treated statistically. As with wing length, the larger 
discal eyespot size of male and female walterorum are significantly 
greater than those of mendocino (P < .05). Analysis indicated that 
only 10 percent of the difference in eyespot size is attributed to the 
difference in wing length between the two species. Considering the 
few characters available, a hybrid index was not deemed neces- 
sary. In terms of qualitative differences, all male walterorum have 
a distinct white apical patch about 2 mm long, on both surfaces 
of the forewing. Males of mendocino may have a similar, but smaller 
white apical patch on the dorsal surface only, thus the presence or 
absence of the white apical patch on the ventral surface is diagnostic. 
Finally, all female walterorum have a bold submarginal black band 
on both the dorsal and ventral surface of the forewing, which is lack- 
ing in female mendocino (Table la, h). 

The average egg weight and length of both walterorum and men- 
docino is similar (ca. 0.0036 g, 2.46 mm. Table 2). One way analysis 
of variance combined with a Duncan multiple range was used to 
compare both egg weight and length. Although no difference was 
found between walterorum and mendocino ova, a statistically signif- 
icant difference in average egg weight and length was found between 
F,a ova and both parental species (P < .05) (Table 2a, b, f, g, h). 
female mendocino usually deposit 70 to 80 eggs within 5 or 6 hours 



Volume 35, Number 1 



5 




Fig. la-f. Female Saturnia and hybrids, la, S. mendocino; lb, 6 Fia x 9 men- 
docino; Ic, 6 walterorum x 9 mendocino = F^a; Id, 6 mendocino x 9 walterorum = 
Fii,; le, 6 Fja X 9 walterorum; If, S. walterorum. Black lines represent 10 mm. 



after mating, while walterorum females deposit 100 to 140 eggs. The 
average fertility of each species was near 97 percent. 

Table 1 presents phenotypic data on the hybrids that were pro- 
duced under controlled conditions. The initial mating of a male wal- 
terorum to a female mendocino produced the hybrids, which were 
nearer in size to walterorum than mendocino. Hybrid Fja females 
also had a distinct submarginal band which was not as well developed 
as that of typical walterorum (Table Ic; Figure Ic). The fore wing to 
hindwing eyespot ratio of these hybrids was <1 as in mendocino. The 
Fla hybrids also resembled mendocino in the appearance of the male 
apical spot (Table Ic). When the Fia male was backcrossed to a female 
mendocino, the resulting adults were almost identical to typical men- 
docino, but the male eyespot ratio was equal to that of walterorum 
even though the eyespot size was greatly reduced relative to walter- 
orum. Females lacked the submarginal band, and expressed an eye- 
spot ratio similar to mendocino, although the absolute size of the spots 



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Journal of the Lepidopterists’ Society 




Fig. 2a-h. Male Saturnia and hybrids. 2a, S. mendocino; 2b, S Fja x 9 mendo- 
cino; 2c, S walterorum x 9 mendocino = Fja; 2d, 6 mendocino x 9 walterorum ~ 
Fib; 2e, 6 Fla ^ walterorum; 2f, S. walterorum; 2g, Wild specimen, Cone Peak, 
Monterey Co.; 2h, Wild specimen. La Panza, San Luis Obispo Co. Black lines repre- 
sent 10 mm. 



were larger. The male backcrossed to a female walterorum pro- 
duced progeny which expressed all of the qualitative characters as- 
sociated with walterorum, while the quantitative characters were in- 
termediate (Table le). In both backcrosses of males to the parent 
species, the fertility was 6 to 9 percent below normal (Table 2d, e). 

Unlike Fja males, the Fia females were almost totally sterile, and 
laid about half the normal number of ova. The eggs which these hy- 
brid females produced were small, and of unusual shape and size, 
with an average weight of only 50-65 percent of normal (Table 2f, g, 
h). Dissection revealed some eggs contained dead, partially formed 
larvae, but the majority of the eggs lacked any observable embryonic 
development. Of the 96 eggs resulting from backcrosses to male wal- 
terorum and mendocino, none was fertile. 

The progeny from the reciprocal cross, 6 mendocino x 9 waiter- 
orum, were the F^, hybrids. The F^, adults were similar in size to 
mendocino, and lacked the apical patch in the males, and submarginal 
bands in females. The eyespot size and ratio was intermediate in the 
females, while in the males the eyespot ratio was close to that of 
walterorum. Unlike F,a females, Fj^ females were fertile, producing 
the normal weight and number of eggs (Table 2j). When the Fjb adults 



Volume 35, Number 1 



7 



were selfed, the resulting Fg larvae were sub vital and only seven 
were reared to adults. Of the five females, two emerged with crippled 
wings, two had thinly scaled wings and one appeared normal; the two 
males were normal. 

The fertility and fecundity of the backcross progeny were tested by 
selfing (Table 2k, 1). The ova size and weight were near normal for 
both crosses. The percent hatch was normal for the ova laid by the 
female whose female parent was walterorum, but none of the eggs 
hatched in the backcross with a mendocino parent. This may have 
been the result either of the pair separating prematurely or sterility. 

In the pure stock of mendocino and walterorum the immature lar- 
val phenotypes, though ver>^ similar, are discrete and non-overlap- 
ping. Larvae from any given hybrid cross could express phenotypes 
of either species, as well as any number of intermediate forms. Thus, 
there was no clear case of phenotypic dominance. Mature larvae of 
each species can best be distinguished by differences in setal pattern 
and length (Tuskes, 1976). 



Discussion 

Much of the biological information regarding the life history of the 
New World Saturnia was summarized by Ferguson (1972), but not 
all of the published information available at the time was correct. In 
describing mendocino, Behrens (1896) gave the type locality as “the 
forests of Sequoia Sempervirens, of the Coast range of Mendocino 
County, Cal.’' Thus, Ferguson (1972) contrasted the “moist coniferous 
forest” inhabited by mendocino, to the arid chaparral habitat of wal- 
terorum in southern California, and implied that this ecological dis- 
tinction might be diagnostic. In fact, mendocino occurs in the arid 
Oak-Digger Pine woodland, and chaparral plant communities, where 
it feeds on manzanita, Arctostaphylos spp. (Ericaceae). Though these 
plant communities may occur adjacent to coastal or canyon redwood 
forests, the ecological and climatological differences are severe (Bak- 
ker, 1971; Major, 1977). In addition to manzanita, there is one report 
(Tilden, 1945) of mendocino larvae feeding on Madrone, Arbutus 
menziesii Pursh (Ericaceae), which on occasion is found in drier 
areas along the border of the redwood community, and opens the 
possibility that mendocino may occur there. 

Saturnia mendocino occurs in the western foothills of the Sierra 
Nevada, from Tulare Co., north into the Cascade range of Siskiyou 
Co. (Fig. 3). One specimen has been collected just north of the Cal- 
ifornia border in Jackson Co., Oregon (Tuskes, 1976), and marks the 
northern limit of well defined chaparral communities; whether men- 
docino occurs farther north on Arctostaphylos or possibly Arbutus 



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Journal of the Lepidopterists’ Society 




Fig. 3. The known distribution of S. walterorurn and S. mendocino in California. 



remains to be determined. The southern limits of mendocino in the 
Coast Range have not been adequately determined. Specimens with 
intermediate phenotypes have been collected in Monterey and San 
Luis Obispo Counties, and will be discussed later. Saturnia walter- 
orum is found from Santa Barbara Co. south to San Diego Co. and 
undoubtedly occurs in Baja California. Though walterorurn is not 
reported from Riverside or San Bernardino Counties, suitable habitat 
for it occurs in both counties. In the coastal chaparral community, 
walterorurn is associated with Rhus laurina Nutt, and Rhus integri- 



\'OLUME 35, Number 1 



9 



folia Benth. & Hook. (Anacardiaceae) while above 1300 m the larval 
host is Arctostaphijlos (Tuskes, 1974). 

The flight time for both species is generally from February to April, 
depending on altitude and seasonal differences. Populations of men- 
docino in the Cascade Range, of those of walterorum in the Laguna 
Mts. at 2600 m may not emerge until April or early June. The emer- 
gence of adults appears to be highly synchronized, and occurs during 
the first few days of warm weather following a protracted cool period. 
The pupae of both walterorum and mendocino possess a patch of 
clear integument over the brain which suggests that daylength may 
act as a cue controlling development as has been demonstrated in 
Antheraea polyphemiis and A. perniji (Williams & Adkisson, 1964) 
and Actias (Miyata, 1974). Whether this mechanism in Saturnia ini- 
tiates development in the spring or controls summer diapause, or both 
remains to be determined. 

Sala & Hogue (1958) mention the development of definable adult 
structures, such as wings and legs in the pupa during early autumn 
and state further that no walterorum pupae remained viable longer 
than one year. We have not found the development of the pupa to be 
different from other North American saturniids. In addition, we found 
that both mendocino and walterorum are capable of surviving at least 
two years in the pupal stage. Differences in pupal development and 
the ability to survive more than a year in the pupal stage may be the 
result of different rearing conditions. 

The third American species in this genus, S. albofasciata, is unique 
when compared to the other two. The adults of this species exhibit 
strong sexual dimorphism, and fly during October and November, 
rather than the spring. Male albofasciata fly and mate in the late 
afternoon, while females oviposit within a few hours after sunset. The 
laiwae feed on Ceanothus (Rhamnaceae) and Cercocarpus (Rosa- 
ceae). Saturnia albofasciata occurs in both the Coast Range and the 
foothills of the Sierra Nevada from Lake Co. south to Los Angeles Co. 
(Ferguson, 1972). Recently specimens have been taken near Julian 
and Campo in San Diego Co., and this species probably occurs in 
northern Baja California. Additional details regarding this moth are 
given by Johnson (1938, 1940) and Hogue et al. (1965). 

Comparison of Larvae 

The larvae of walterorum and mendocino are very similar but can 
be distinguished at all stages of development. Color polymorphism 
occurs in both species, and the larvae of mendocino are especially 
variable. The first and last instars of walterorum have been described 
(Sala & Hogue, 1958) and all the developmental stages of mendocino 



10 



Journal of the Lepidopterists’ Society 




Fig. 4a-d. Dorsal and lateral view of Saturnia larvae. Second (4a) and third instar 
(4b) larvae of S. walterorum; second (4c) and third instar (4d) larvae of S. mendocino. 



were described by Comstock (1960), although he did not recognize 
the fourth instar as the final larval stage due to the loss of the brood. 
We provide here a more complete description of variation, and list 
diagnostic characters for both species. 

Figure 4 illustrates the dorsal and lateral view of the second and 
third instar larvae of walterorum and mendocino. The setae were 
omitted to emphasize pattern differences. In second instar waltero- 
rum, the dorsal scoli of abdominal segments I and VIII are enclosed 
in posterior and anterior dorsal black bands, while in mendocino 
these two dorsal bands enclose the dorsal scoli of the meso- and meta- 
thoracic segments as well as those of abdominal segments I, II, VIII 
and IX. The meso- and metathoracic dorsal scoli of mendocino are 
also smaller than those of walterorum. 

In the third instar, walterorum lacks the posterior and anterior dor- 
sal bands of abdominal segments I and VIII, which are present in 
mendocino. The length of the lateral stripe is variable in mendocino, 
and may lie as illustrated, or may connect with the thoracic transverse 
l)and; in extreme cases a connection also exists with the caudal band, 
forming a rectangle, enclosing the dorsal abdominal scoli. Upon molt- 
ing into the tliird instar, the ground color of mendocino is yellowish 
pink dorsally and dull, salmon pink laterally. The day after ecdysis a 








Volume 35, Number 1 



11 



rapid color change occurs; some larvae become lemon yellow, others 
turn light green, while a smaller number are a salmon pink with a 
yellow tinge. The green phase of both species is probably the most 
common. A similar polymorphism occurs in walterorurn except that 
the yellow phase usually has an orange tint. 

The mature larvae of walterorurn and mendocino are very similar 
and can be reliably separated only by the greater number of dark 
proleg setae in mendocino (Tuskes, 1976). Ferguson (1972) noted that 
the description by Sala & Hogue (1958) of the mature walterorurn 
larvae made no mention of a prominent yellow lateral line seen in 
mendocino and inferred that this was a means to distinguish the two 
species. This line does exist in walterorurn, and extends from the 
caudal area to the metathoracic segment, passing just below the spi- 
racles. Donahue (1979) illustrated in eolor a mature walterorurn larva 
photographed by one of the authors. We have also found that unlike 
mendocino larvae, which pass through only four instars, approxi- 
mately 62 percent of the walterorurn have five instars, the remainder 
matured in four. Furthermore, about 80 percent of the larvae with 
five instars developed into females. Fourth and fifth instar larvae are 
identical in color and pattern and as in the third instar, the green color 
phase is the most common. In mendocino, lemon yellow larvae often 
have a greenish tint, while walterorurn larvae in this phase are yel- 
lowish-orange. The third color form is more difficult to describe but 
has been called mauve by Comstock (1960) and salmon by Sala & 
Hogue (1958). We have noted Yolo Co. mendocino in this phase are 
a salmon-pink with a yellowish cast. Infrequently a variation occurs 
which is a richer color, close to a reddish-brown. Williams (1905) 
reeorded the green form and this brownish phase of mendocino from 
Mt. Shasta, and Tilden (1945) collected this color form in Santa Cruz 
Co. Interestingly, the color of the third instar larva cannot be used 
to predict the color phase of the mature larva. 

Hybridization Studies 

We have directed the results of our hybridization studies toward 
three areas of investigation: a functional measure of genetic incom- 
patibility between walterorurn and mendocino through all stages of 
development; an evaluation of potential barriers to interspecific mat- 
ing; and a comparison of hybrid phenotypes to specimens from geo- 
graphically intermediate populations. 

The results indicate a certain degree of genetie compatibility be- 
tween walterorurn and mendocino. The successful production of Fj 
adults, using either species as the female parent, indicates that the 
mode of gene expression throughout development is compatible in 



12 



Journal of the Lepidopterists' Society 



the two species, and indirectly implies a great deal of allelic homol- 
ogy. By contrast, developmental incompatibility has been demonstrat- 
ed in hybrids of closely related species of Callosamia (Peigler, 1977) 
and Phyciodes (Oliver, 1978). During the formation of gametes in the 
Fj progeny, the once equal representation of chromosomes from each 
parental species is randomly assorted in meiosis. Dissimilarities in 
chromosomes can cause aborted gonad or gamete formation especially 
in the heterogametic sex (Dobzhansky, 1970). Thus, sterile hybrid 
females are common in studies with Hyalophora (Sweadner, 1937; 
Collins, 1973), and European Saturnia (Standfuss, 1900, 1901a, b). 
Yet in this study fertile Fj hybrid females were produced when the 
female parent was walterorum, while the reciprocal cross produced 
sterile females which possessed about half of the normal number of 
ova. These results suggest that perhaps the observed nonreciprocal 
sterility is somehow linked to the genetic basis of sexual dimorphism, 
which is strongly expressed in walterorum but less so in mendocino. 
In Lepidoptera ZW represents the heterogametic female sex chro- 
mosome combination, and ZZ denotes the male sex chromosomes. 
Perhaps the action in hybrid females of the Z chromosome from wal- 
terorum is incompatible with that part of the W chromosome from 
mendocino which affects gametogenesis. In male hybrids of various 
combinations the eyespot size is close to walterorum^ while in female 
hybrids the eyespots grade from small to large, depending on the 
parentage of the cross (Table 1, Fig. 1). This is additional circumstan- 
tial evidence of disruptive effects caused by the Z chromosome of 
walterorum. Further, it appears that at least some of the sexual di- 
morphism observed in female walterorum (ground color and sub- 
marginal forewing band) is sex limited and is not carried by alleles 
on the W chromosome. Though none of these characters are expressed 
by male walterorum, they are transmitted by the male to the Fj fe- 
male progeny when mated to a female mendocino. In addition, it 
would also appear that discal eyespot size and ground color are poly- 
genic and not expressed as simple dominant or recessive traits. 

Although F,a females were sterile, F^, females backcrossed to the 
parent species had nearly normal fertility and fecundity. In backcross- 
es, partial genome integrity is preserved via the non-hybrid parent. 
By contrast, F 2 females were sterile. High mortality in the F 2 gener- 
ation may be ascribed to hybrid breakdown, the disruption of highly 
integrated parts of the genome (Dobzhansky, 1970, 1977). 

Analysis of Intermediate Populations 

Populations of Saturnia exhibiting intermediate characters were 
found in the central Coast Range (Fig. 3). Three male Saturnia were 



Volume 35, Number 1 



13 



trapped, using an Fj female as bait, near Cone Peak, Santa Lucia 
Mountains, Monterey Co. These specimens are the size of men- 
docino but possess 26 percent larger hindwing eyespots (Fig. 2g). The 
ventral apical mark appears as a trace, similar to many Fj male spec- 
imens. Ferguson (1972) and Tuskes (1974) cite Tilden's capture of 
three male walterorum in the La Panza Range, San Luis Obispo 
Co. but upon examining these specimens we found that they are 
not typical walterorum. We collected three additional males which 
were attracted to a female mendocino near La Panza Summit. The 
La Panza males have larger eyespots than the Cone Peak specimens 
and exhibit the white apical mark of walterorum. They resemble 
mendocino in size and in having a larger hindwing eyespot than fore- 
wing eyespot, as do the Cone Peak males (Fig. 2). If the eyespot size 
relative to forewing length of these intermediate specimens is com- 
pared to the Santa Barbara walterorum, we find that the hindwing 
eyespot of the Cone Peak males, and fore- and hindwing eyespots of 
La Panza males are proportionately larger. As mentioned in Results, 
difference in overall size accounts for only 10 percent of the differ- 
ence in eyespot size between typical mendocino and walterorurn. 
Thus, both the Cone Peak and La Panza Saturnia appear like men- 
docino in overall size and in having an eyespot ratio less than one, 
but have prominent apical marks and larger eyespots; the walterorum 
characters are more pronounced in the more southern La Panza pop- 
ulation. 

One of the La Panza males was mated to the female mendocino 
and the resulting larvae exhibited mixed larval phenotypes. Five fe- 
males and one male were reared to maturity; the females were the 
size of mendocino and lacked any trace of a submarginal black fore- 
wing band, but showed the ground color of walterorum, much like 
the laboratory F^ hybrids. 

We can offer only tentative conclusions about the taxonomic status 
of the Saturnia in the Santa Lucia and La Panza mountains. A larger 
sample needs to be collected, especially of the more diagnostic fe- 
males. However, the available phenotypic evidence, combined with 
the demonstrated lack of reproductive barriers, and high degree of 
genetic compatibility in hybrids, suggests that at some time in the 
past the Saturnia in the central California Coast Range could have 
undergone a period of hybridization and introgression between men- 
docino-like and walteroru7n-\ike populations. We theorize below that 
this event may have been secondary to the divergence of these taxa, 
perhaps during the post Pleistocene xerothermic event. 

In summary, Saturnia mendocino and S. walterorum can best be 
classified as semispecies, as exemplified by Drosophila paulistorum 



14 



Journal of the Lepidopterists’ Society 



(Dobzhansky et al., 1977). Morphological differences between the two 
taxa are slight; mature larvae, cocoons, pupae, and adult males are 
very similar, while immature larvae and adult females are distinct. 
No prezygotic barriers to reproduction exist under laboratory condi- 
tions. Postzygotic mechanisms act to reduce reproductive fitness in 
certain primary crosses, but backcrosses can be fertile in both sexes. 
The F., adults are sterile and frequently malformed. The historic iso- 
lation between mendocino and walterorum in the southern Coast 
Range has probably been topographic. Between the southern Sierra 
Nevada and the Coast Range there are expanses of desert and nu- 
merous small mountain ranges which lack host plants of either 
species. In the central Coast Range, Arctostaphylos chaparral is dis- 
continuous, separated by other types of vegetation and intervening 
lowlands (Hanes, 1977). Such discontinuities appear to exist in north- 
ern Santa Barbara and southern San Luis Obispo counties, and may 
represent the boundary between the two species. 

Phylogeny of Saturnia and Agapema 

Recent phylogenies of Lepidoptera have combined morphological 
and biogeographical data with a comparative knowledge of foodplant 
preferences (Ehrlich and Raven, 1965), based on the general finding 
that host plant choices are taxonomically specific and evolutionarily 
conservative. Conversely, evolutionary radiation is often accom- 
panied by new host plant associations. Such insect-plant relationships 
are thought to represent coevolution at the community level (Whit- 
taker & Feeny, 1971; Feeny, 1973). 

The American Saturnia and the closely related genus Agapema 
seem to represent examples of organisms coevolving with the sclero- 
phyllous members of the Madro-Tertiary flora in western North Amer- 
ica. While fossils of these moths are lacking, fossil records of their 
present day host plants do exist, and knowledge of floral distribution 
through time provides a framework for a phylogenetic discussion. We 
must assume that the present day host plants of Saturnia and Aga- 
pema reflect ancient associations, at least at the family level. Before 
reviewing the fossil flora evidence, we discuss our reasons for in- 
cluding Agapema in the discussion and briefly compare the host 
plants of Eurasian Saturnia and their allies with their North American 
relatives. 

The genus Agapema is morphologically distinct from Saturnia but 
is closely related to it; Ferguson (1972) separates these genera but 
Michener (1952) treated Agapema as a subgenus of Saturnia. Many 
European and Asian Saturnia, as well as related genera such as Dic- 
tyoploca and Caligula are similar to Agapema in pattern and color- 



Volume 35, Number 1 



15 



ation and are also sexually monomorphic nocturnal fliers. The larvae 
of Dictyoploca and Caligula are adorned with long hairs and bear a 
resemblance to the larvae of Agapema in this respect. As noted by 
Hogue et al. (1965) the nocturnal female of S. albofasciata somewhat 
resembles the nocturnal gray-colored adults of Agapema. Saturnia 
albofasciata also resembles the European S. pavonia in flight rhythm 
and sexual dimorphism, and an ancestral link between these two 
species has been suggested by Hogue et al. (1965). Yet, Ferguson 
(1972) notes the genitalia of Agapema are more primitive and quite 
similar to S. pavonia, while the three American Saturnia, especially 
albofasciata, are more specialized and divergent from the European 
Saturnia. Lemaire (1979) also stresses the uniqueness of the Ameri- 
can Saturnia (which he places in the subgenus Calosaturnia) and 
within this group he further recognizes S. albofasciata as the most 
divergent member, even though this species resembles Old World 
species in retaining the aedeagus, which is lost in S. walterorum and 
S. mendocino. Thus the similarities between S. pavonia and S. al- 
bofasciata may be parallelisms; such phenotypic and phonological 
flexibility is characteristic of Saturniidae in general. 

The Old World Saturnia, as well as the related Eurasian genera 
Caligula, Cricula, and Dictyoploca, tend to be polyphagous; impor- 
tant host plant families include Ericaceae, Rosaceae, Salicaceae, and 
Fagaceae, but not, apparently, Rhamnaceae. Several trends are ap- 
parent in comparisons of New and Old World host plants. The eri- 
caceous preference of S. pavonia is seen in S. walterorum and S. 
mendocino, but not in the superficially similar S. albofasciata, thus 
further substantiating a more derived rather than ancestral status for 
this species. Saturnia albofasciata does retain a widespread Old 
World preference for rosaceous plants in its inclusion of Cercocarpus 
as a host plant, although it is possible this plant merely resembles 
Rhamnaceae biochemically and that this is the basis for its utilization 
by the moth. Rhamnaceae may be a new host plant group acquired 
during the New World evolution of Saturnia and Agapema. The lar- 
vae of A. homogena feed on Rhamnus in Arizona (Mr. Kenneth Han- 
sen, pers. comm.) and are said to refuse Arctostaphylos in captivity. 
Other species of Agapema feed principally on Condalia and related 
Rhamnaceae. As mentioned above, S. albofasciata feeds on Ceano- 
thus (Rhamnaceae). Thus, morphological similarity establishes a tie 
between Agapema and Eurasian Saturnia and related genera, and 
general morphology and host plant selection provides a link between 
Agapema and American Saturnia, especially albofasciata. Rhus laur- 
ina and R. integrifolia are food plants only of walterorum and may 
be associated with this species’ divergence from mendocino, as dis- 



16 



Journal of the Lepidopterists’ Society 



Table 3. Fossil records of present day Saturnia-Agaperna host plant genera. 



Miocene 


Pliocene 


Techachapi; Southern Calif. 
(Axelrod, 1939): 
Arctostaphylos glandulosa 
Cercocarpus betuloides 
Rhamnus calif ornica 
Rhus integrifolia 
Ceanothus cuneatus 


Anaverde, Mt. Eden, Pirn George; 
Southern Calif. (Axelrod, 1950): 
Arctostaphylos 
Cercocarpus 
Rhamnus 
Rhus laurina 


Aldrich-Fallon-Middlegate; 

Interior Nevada (Axelrod, 1956): 
Arbutus 
Ceanothus 
Cercocarpus 


Table Mountain, Remington Hills, 
Chalk Hills; Cent. Calif 
(Chaney, 1944): 

Arbutus 

Ceanothus cuneatus 
Cercocarpus 
Arctostaphylos 
Rhamnus 


Mint Canyon; Southern Calif. 
(Axelrod, 1940): 
Ceanothus cuneatus 
Cercocarpus betuloides 
Rhamnus crocea 


Mulholland; Coastal Central Calif. 
(Axelrod, 1944): 

Ceanothus 
Cercocarpus 
Rhamnus 
Arbutus 
Rhus laurina 



cussed below. In summary, no American species in either genus pos- 
sesses both primitive Old World genitalic structure and host plant 
preferences. It is our thesis that these New World genera became 
specialized by coevolving with their host plants as climate and to- 
pography changed during the late Tertiary and Quaternary. 

The evolution of sclerophyllous plants, as part of the Madro-Ter- 
tiary flora, with which the Saturnia are closely associated, has been 
dealt with in detail by Axelrod and others (Table 3). Axelrod (1977) 
presents a summary discussion and we cite other original papers. His 
thesis rests on the premise that ancient climates can be deduced from 
the species composition of geofloras, whose leaf shapes and structures 
are clues to their ecological requirements. In many cases these an- 
cient species closely resemble living forms. As climates and topog- 
raphy changed, the distribution of plant species shifted accordingly. 
Those groups preadapted to xeric conditions underwent rapid specia- 
tion (e.g., Ceanothus, Arctostaphylos, Quercus), while plants depen- 
dent on summer rain were displaced as the climate became cooler 
and drier. Thus, during the Miocene-Pliocene there was a general 
coastward movement of Madro-Tertiary flora. 

The Madro-Tertiary flora in the early Tertiary developed as more 



Volume 35, Number 1 



17 



xeric tolerant elements of a very generalized, diverse woodland, in- 
cluding deciduous species, which enjoyed a moderate climate of sum- 
mer rains and mild winters. Due to the lack of major topographic 
relief and the widespread floras, it is possible that in the Miocene the 
Saturnia-Agapema ancestral stock existed as one or a few distinct 
species, having arrived from Asia via a land bridge during Eocene- 
Oligocene times. Yet, modern host plant relationships could have 
evolved at this time. 

At the time of the Pliocene the various genera of food plants utilized 
by Saturnia and Agapema were all members of a single community 
which extended as a more or less continuous flora throughout the 
present day Great Basin and Southwest. Border redwood, redwood, 
and chaparral communities occurred to the north in the Sierra, then 
only 1000-1300 m in elevation. The middle Pliocene was probably 
the last period when central and southern California coastal floras 
were intermixed (Chaney, 1944). We can surmise that Saturnia had 
not necessarily diverged into the precursor populations of mendocino 
and walterorum since the distribution of Arctostaphylos was so wide- 
spread. Perhaps the more northern populations also fed on Arbutus 
as members of a tan oak-madrone-canyon oak community, while 
southern populations extended their ecological tolerance into a warm- 
er coastal community containing Rhus. 

Climatic and geological factors in the late Pliocene and Pleistocene 
caused segregation of separate plant communities from more gener- 
alized communities. More extreme seasonal fluctuations developed 
and in general the climate was becoming cooler and drier. The con- 
tinuing rise of the Sierra Nevada and the subsequent uplift of the 
Santa Ana and San Gabriel Mountains as well as the Coast Ranges 
occurred at this time. These altitudinal changes and the accompany- 
ing rain shadows produced dramatic environmental dines. Chaparral 
as a distinct and widespread community type probably first appeared 
in the Pleistocene as an altitudinal segregate on the lower slopes of 
uplifting mountains. The final disappearance of summer rains gave 
rise to a Mediterranean climate along the coast but the interior pen- 
etration of the moderating coastal climate was eliminated as mountain 
building occurred. We can hypothesize that as the southern California 
Rhus-Arctostaphylos association began to separate into montane and 
coastal communities the moths expanded their distribution accord- 
ingly. Northern California populations diverged into mendocino on 
manzanita in the Coast Range and in the Sierras, separated by an 
increasingly inhospitable valley. A northern bridge of Arctostaphylos 
and perhaps Arbutus in the Sierra-Cascades provided genetic conti- 
nuity to this wide ranging species. 



18 



Journal of the Lepidopterists’ Society 



Saturnia albofasciata probably arose as a separate entity in south- 
ern California in association with the more xeric-adapted Ceanothus 
cuneatus and Cercocarpus betuloides. Its appearance in the Coast 
Ranges would then be one of invasion as C. cuneatus and C. betu- 
loides spread on the uplifting coastal mountains. Thus, the sympatry 
of mendocino and albofasciata may be rather recent. 

The rain shadow of the Sierras and southern California mountains 
produced an intervening desert which isolated the Arizona deriva- 
tives of Saturnia. Agapema homogena may be a relict species as it 
now inhabits montane areas of summer rains (Colorado, Arizona, New 
Mexico, west Texas and portions of northern Mexico) and emerges in 
early summer. The desert species of Agapema probably appeared at 
this time although it is possible that they speciated earlier in Mexico 
and subsequently invaded the developing American deserts to the 
north. 

The phenomenon of the xerothermic period may explain the ap- 
parently intermediate populations of Saturnia in the Santa Lucia and 
La Panza mountains. The xerothermic of 3000 to 8500 years ago was 
a sudden warming period between the last glaciation and the more 
recent cooling period (Axelrod, 1966). This change in climate appears 
to have forced chaparral species such as Arctostaphylos glauca Lindl. 
north into the Coast Ranges such that relict populations now exist as 
far north as Mt. Hamilton. Similarly, Rhus laurina and R. integrifolia 
have an oddly disjunct population near Cayucos, 130 km N of their 
normal range. The northern movement of all these plants during a 
brief period of warmth may have produced a temporary event of hy- 
bridization and introgression between S. mendocino and S. walter- 
orum. 

We can hypothesize that the Saturnia responded to the same en- 
vironmental changes as their host plants and evolved phenological 
and developmental adaptations. The highly synchronized spring adult 
emergence and facultative egg development of mendocino and wal- 
terorum allow the larvae to exploit the early growth of their food 
plants. The egg of albofasciata represents an alternative modification, 
allowing the larva to overwinter and emerge in the early spring to 
feed on new growth. Perhaps the fall flight of this species is a direct 
result of this adaptation. The genetic potential for this adaptation may 
well be ancestral; an overwintering egg occurs in the Asian Caligula 
and Dictyoploca, and in Arizona populations of Agapema galbina. 
The pupae of walterorurn and mendocino pass through the hot dry 
summer months, as well as the winter. The open mesh cocoon con- 
struction is especially well developed in the desert species of Aga- 
pema, and much less so in the montane A. homogena. The loose mesh 
cocoon may aid in ventilation, keeping the pupa cooler. 



Volume 35, Number 1 



19 



Hogue et al. (1965) proposed that both mendocino-walterorum and 
the genus Agapema arose sympatrically from an albofasciata-\ike 
ancestor by means of dual mutations affecting coloration and flight 
times, such that dull colored night flying mutant males would en- 
counter more similarly colored nocturnal females, and brightly col- 
ored mutant diurnal females would encounter normal brightly colored 
diurnal males. In this way arose the brightly colored diurnal walter- 
orum-mendocino line, which resembles male albofasciata, and the 
dull colored nocturnal Agapema, which are similar to the female of 
albofasciata. It should be pointed out that female flight occurs only 
after mating, and that pheromones, not chance encounters in flight, 
control mating response. Furthermore, Saturniidae are poor candi- 
dates for sympatric speciation as judged by the criteria of current 
models (Wilson et ah, 1975). They are present in large mobile, more 
or less randomly mating populations, in which the uniting of rare 
mutants is unlikely. Since the pheromone system controls mating, 
mutants with allochronic mating behavior would be severely selected 
against. Rather, we feel that the Saturnia- Agapema phylogeny is one 
of coevolution with Madro-Tertiary flora in which moths and their 
sclerophyllous host plants adapted to changing climate, primarily by 
altering developmental phenomena. The continual lateral and alti- 
tudinal redistribution of plant communities, especially during the 
Pliocene-Pleistocene, provided ample opportunity for allopatric spe- 
ciation to occur. In this context the West Coast Saturnia are important 
examples of endemic species which evolved in response to the ap- 
pearance of a Mediterranean climate. 

Acknowledgments 

We wish to thank Arthur Shapiro for reviewing the original manuscript and Steve 
McElfresh for the loan of specimens. The California Insect Survey, Dept, of Entomol- 
ogy, Univ. Calif. Berkeley provided the map used in Fig. 3. 



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1944. The Mulholland flora. Carnegie Inst. Wash. Pub. 533: 103-146. 

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590: 119-158. 

1956. Mio-Pliocene floras from west-central Nevada. Univ. Calif. Pub. Geol. 

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1966. The early Pleistocene Soboba flora of Southern California. Univ. Calif. 

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20 



Journal of the Lepidopterists’ Society 



1977. In Barbour, M., & J. Major, eds., Terrestrial vegetation of California. 

Wiley, New York. 1002 pp. 

Barker, E. 1971. An island called California. Univ. Calif. Press, Berkeley. 357 pp. 
Behrens, J. 1876. Description of a new saturnian. Can. Entomol. 8: 149. 

Chaney, R. 1944. Pliocene floras of California and Oregon. Carnegie Inst. Wash. Pub. 
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Comstock, J. A. 1960. Life history notes on a saturniid and two lasiocampid moths 
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Collins, M. M. 1973. Notes on the taxonomic status of Hyalophora Columbia (Sa- 
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Dobzhansky, T. 1970. Genetics of the evolutionary process. Columbia Univ. Press, 
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Dobzhansky, T., F. Ayala, G. L. Stebbins & J. W. Valentine. 1977. Evolution. 
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Donahue, J. P. 1979. Strategies for survival, the cause of a caterpillar. Terra 17: 3-9. 
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Hogue, C. L., F. P. Sala, N. McFarland & C. Henne. 1965. Systematics and life 
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1940. The allotype of Calosaturnia albofasciata (Lepidoptera, Satumiidae). 

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272-276. 



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Whittaker, R. & P. Feexy. 1971. Allelochemics: chemical interactions between 
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(U.S.A.) 72: 5061-5065. 



Journal of the Lepidopterists" Society 
35(1), 1981, 21 



RECENT ADDITIONS TO THE COLLECTION OF THE AxMERICAN MUSEUM 

OF NATURAL HISTORY 

Dr. Cyril F. dos Passos has donated his collection of 65,382 butterflies to the Amer- 
ican Museum of Natural History. Of this total, 64,052 specimens are mounted and 
identified; 57,870 are from North America and 6182 are from Europe; 1330 are un- 
mounted or unidentified. Included in the collection are 464 paratypes (no holotypes 
or allotypes) and 617 slides (124 venation, 493 genitalia). Dr. dos Passos started build- 
ing his collection in 1929; it undoubtedly represents the single largest and most com- 
plete one of North American butterflies ever made by one individual. It far surpasses 
the two previous large collections of butterflies (no moths) received by the Department 
of Entomology, namely those of J. D. Gunder (27,000 North American specimens, 
received in 1937) and V. G. L. van Someren (23,000 African specimens, received in 
1970). The addition of this collection gives the American Museum of Natural History 
an unrivaled collection of North American butterflies. 

The museum has also received the collection of the late Bernard Heineman, con- 
sisting of 7075 mounted butterflies and moths. Of these, 2857 were from Jamaica, with 
the great majority being butterflies. This is the largest private collection of Jamaican 
butterflies ever made, and it served as the starting point for the 1972 book entitled, 
“Jamaica and its butterflies” by F. Martin Brown and Bernard Heineman. The other 
4218 specimens represent a world- wide collection made by Mr. and Mrs. Heineman 
on their various trips throughout the world. 

Frederick H. Rixdge, Department of Entomology, The American Museum of Nat- 
ural History, New York, New York 10024.